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研究生:黃青萸
研究生(外文):Ching-Yu Huang
論文名稱:南仁山森林落葉層兩棲類與無脊椎動物對落葉分解之影響
論文名稱(外文):Effects of the litter amphibians and the litter invertebrates on litter decomposition in Nanjenshan forest
指導教授:侯平君侯平君引用關係
指導教授(外文):Ping-Chun Lucy Hou
學位類別:碩士
校院名稱:國立成功大學
系所名稱:生物學系碩博士班
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2002
畢業學年度:90
語文別:英文
論文頁數:78
中文關鍵詞:落葉分解無脊椎動物群落兩棲類捕食效應分解常數
外文關鍵詞:decay ratepredationAmphibianinvertebrate communitylitter decomposition
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摘要
在陸域生態系統中,落葉分解是重要的生態過程。根據前人的研究已知落葉分解主要受落葉品質、氣候因子及土壤生物相所影響。最近的文獻指出:熱帶地區的土壤動物對於落葉分解具有重要的貢獻。此外,森林底層兩棲類也可能透過捕食大型土壤動物,間接影響落葉分解速率。本研究主要探討位處台灣南部的南仁山低地雨林,森林落葉層兩棲類、無脊椎動物及落葉分解速率三者間的關係。實驗於南仁山生態保護區之古湖樣區進行。論文分為三個章節,第一章探討不同網目(0.1 mm、1 mm、3.5 mm)之枝葉包中,無脊椎動物群落和江某(Schefflera octophylla)葉片分解速率之間的關係。第二章為森林底層兩棲類密度及群落調查,用於第三章實驗之參考。第三章則以野外圍籬實驗,探討盤谷蟾蜍 (Bufo bankorensis)的捕食效應,對無脊椎動物群落及落葉分解速率的影響。結果發現,大網目(3.5 mm)的枝葉包中,落葉分解速率最快且有較多的蜘蛛、等足目及倍足綱等的大型無脊椎動物。小網目枝葉包中,彈尾目動物數量明顯較中、大網目枝葉包高,可能為小網目枝葉包中缺乏捕食者(尤其是蜘蛛)所造成。此外,落葉分解速率與無脊椎動物密度有顯著的相關性。南仁山森林底層兩棲類密度雖較美洲熱帶森林低,但卻高於其他亞洲熱帶森林。背風之溪谷型植被區的兩棲類密度為每100平方公尺10.2隻,較迎風之緩風坡型植被區每100平方公尺3.5隻高,且兩區中小蛙所佔的比例均較成蛙高。在圍籬實驗部分,盤谷蟾蜍的存在雖然並未改變無脊椎動物群落之組成,但卻可減少枝葉包中無脊椎動物的密度。然而,盤谷蟾蜍的存在對落葉分解速率未有顯著的影響。上述結果顯示,在南仁山亞熱帶森林中,落葉大型無脊椎動物可以調節彈尾目族群的數量,而無脊椎動物的密度為預測落葉分解速率的一個良好指標。雖然圍籬的設置可能減少兩棲類的捕食,但結果仍顯示兩棲類的捕食有調控落葉層無脊椎動物密度的潛力,然而是否能間接影響落葉分解速率,需更進一步的研究。
Abstract
Litter decomposition is an important process in terrestrial ecosystems. Previous studies have demonstrated that decomposition is influenced by three factors: litter quality, climate, and soil biota. Recent studies pointed out that soil fauna play an important role in litter decomposition in the tropical forests. In addition, litter amphibians may also influence the rate of decomposition indirectly by feeding on litter macro-invertebrates. The purpose of this study is to examine the relationships among litter decomposition, litter invertebrates, and litter amphibians in a lowland rain forest of southern Taiwan. The experiments were carried out in Guhu plot in Nanjenshan Nature Reserve and are presented in three chapters. In chapter one, I investigated the relationship between invertebrate community and decomposition rate of Schefflera octophylla leaves in litterbags of different mesh sizes. In chapter two, I examined densities and composition of litter amphibians in Nanjenshan forest for experimental design in the following chapter. In chapter three, I examined the predatory effect of Bufo bankorensis on litter invertebrates and decomposition rates in the field enclosures. I found the rate of litter decomposition and the abundance of macro-invertebrates were higher in the coarse-mesh litterbags. The higher abundance of collembolans in the fine-mesh litterbags might result from the lack of predatory macro-invertebrates in the bags. In addition, there is a close relationship between the rate of decomposition and the total faunal density. Densities of the litter amphibian in Nanjenshan forest were higher than those in other Asian tropical forests and were lower than those in Neotropical forests. Mean density of litter amphibians in the creek site (10.2 individuals per 100 m2) was higher than that in the windward site (3.5 individuals per 100 m2). The litter amphibians consist of a large proportion of juveniles. The presence of Bufo bankorensis did not significantly affect the composition of invertebrate, however, it did slightly reduce the total density of invertebrates in the litterbags. Nevertheless, the presence of Bufo bankorensis had no significant effect on the rate of decomposition. The results suggest that predatory macro-invertebrates can regulate the abundance of collembolans and the total density of litter invertebrates is a good indicator for predicting the rate of decomposition. Although the enclosures may interfere with the feeding of the toads, the results indicate that predation of litter amphibians have a potential effect on the density of litter invertebrates. However, the indirect effect of amphibian on rate of litter decomposition in tropical forests needs further studies.
TABLE OF CONTENT Page
CHINESE ABSTRACT ................................................................................... I
ENGLISH ABSTRACT ................................................................................ II ACKNOWLEDGEMENT ............................................................................. IV
LIST OF TABLES ....................……………................................................. VIII
LIST OF FIGURES ……………................................................................... X
GENERAL INTRODUCTION ..................................................................... 1
Chapter
1. Invertebrate community and litter decomposition in a lowland rain forest
of Taiwan: effects of litterbag mesh size ....................................... 3
Introduction ........................................................................................... 3
Materials and Methods .......................................................................... 5
Studying area ................................................................................. 5
Experimental design ...................................................................... 6
Litterbags ....................................................................................... 6
Fauna extraction and identification ............................................... 7
Rates of decomposition ................................................................. 7
Statistical analysis ......................................................................... 8
Results ................................................................................................... 9
Invertebrate communities in litterbags of different mesh sizes ....... 9
Rates of litter decomposition .............................................................. 11
Relationships between litter fauna and decomposition ...................... 11
Discussion ................................................................................................. 12
Faunal density and composition in the litterbags ............................... 12
Rates of mass loss at different stages ................................................ 13
Relationships between decay rates and faunal density and diversity ...... 13
2. Densities and composition of the litter amphibians in Nanjenshan
subtropical forest ............................................................................. 15
Introduction ............................................................................................. 15
Materials and Methods ............................................................................ 17
Studying area ..................................................................................... 17
Experimental design .......................................................................... 18
Quadrat sampling ............................................................................... 18
Statistical analysis .............................................................................. 18
Results ...................................................................................................... 20
Environmental variables ..................................................................... 20
Density and composition of the litter amphibians .............................. 20
The abundance of juveniles ................................................................ 21
Discussion ................................................................................................. 22
Litter amphibians in Nanjenshan forest .............................................. 22
Litter amphibians in the two sites ....................................................... 23
The juvenile anurans ........................................................................... 24
3. Effects of amphibian predation on litter invertebrates and rates of decomposition ..................................................................................... 25
Introduction ............................................................................................... 25
Materials and Methods .............................................................................. 27
Studying area ....................................................................................... 27
Experimental design ............................................................................ 28
Litterbags ............................................................................................. 29
Fauna extraction and identification .................................................... 29
Rate of decomposition ........................................................................ 30
Statistical analysis .............................................................................. 30
Results ...................................................................................................... 31
Effects of enclosures .......................................................................... 31
Amphibians ........................................................................................ 31
Litter invertebrate communities ......................................................... 31
Rates of litter decomposition ............................................................. 32
Discussion ................................................................................................ 34
The effects of enclosures ................................................................... 34
The effects of litterbags ..................................................................... 34
The effects of treatment ..................................................................... 35
Decomposition rates .......................................................................... 35
CONCLUSION ............................................................................................... 37
BIBLIOGRAPHY ........................................................................................... 38

LIST OF TABLES
Table Page
1.1 Analysis of variance for the total invertebrate densities in litterbags in Nanjenshan forest, Taiwan .................................................................. 44
1.2 One-Way analysis of similarity for invertebrate community in the litterbags of between the creek and the windward sites ................................... 45
1.3 Percentage composition, diversity and density of invertebrate community in the coarse-mesh litterbags from Nanjenshan forest, Taiwan ........... 46
1.4 Percentage composition, diversity and density of invertebrate community in the medium-mesh litterbags from Nanjenshan forest, Taiwan ........ 48
1.5 Percentage composition, diversity and density of invertebrate community in the fine-mesh litterbags from Nanjenshan forest, Taiwan ............... 50
1.6 Analysis of variance for the decomposition of Schefflera octophylla in Nanjenshan forest, Taiwan ............................................................... 52
1.7 Mean annual decay rates (k) of Schefflera octophylla leaves in litterbags of different mesh sizes in two sites over a period of 124 days in the field ................................................................................................ 53
1.8 Relationships between the mean densities and diversities of total invertebrates and the annual decay rates (k) of Schefflera octophylla in Nanjenshan forest, Taiwan ................................................................................ 54
2.1 Species richness, total densities, and diversities of litter amphibians in two sites of Nanjenshan forest, Taiwan in 2001 ............................ 55
2.2 Density of the litter amphibians in two sites of Nanjenshan forest, Taiwan in 2001 ........................................................................................... 56
3.1 The environmental variables inside and outside the enclosures during the experimental period in Nanjenshan forest, Taiwan ....................... 57
3.2 Effect of the experimental enclosures on the initial densities of litter invertebrate in Nanjenshan forest, Taiwan .................................... 58
3.3 Analysis of variance for the total invertebrate densities in litterbags from Nanjenshan forest, Taiwan ............................................................. 59
3.4 Invertebrate densities in the litterbags during the experiment and in the litter samples within the enclosures at the end of experiment ................ 60
3.5 Invertebrate composition, diversity and density in the litterbags at six sampling times from the control and the amphibian-excluded enclosures ....................................................................................... 61
3.6 Analysis of variance for mass remaining in the litterbags containing fallen leaves of Schefflera octophyll from the control and amphibian- excluded (treatment) enclosures in Nanjenshan forest, Taiwan ..... 63

LIST OF FIGURES
Figure Page

1.1 Total density of invertebrates in the litterbags from Nanjenshan forest, Taiwan .......................................................................................... 64
1.2 Total number of invertebrates in the litterbags from Nanjenshan forest, Taiwan .......................................................................................... 65
1.3 Abundance of collembolans in the litterbags from Nanjenshan forest, Taiwan .......................................................................................... 66
1.4 Abundance of predaceous invertebrates in the litterbags from Nanjenshan forest, Taiwan ............................................................................... 67
1.5 Abundance of saprophagous invertebrates in the litterbags from Nanjenshan forest, Taiwan ........................................................... 68
1.6 Abundance of Acari in the litterbags from Nanjenshan forest, Taiwan .......................................................................................... 69
1.7 Multi-dimensional scaling (MDS) plot of invertebrate communities in the litterbags from the creek and the windward sites ......................... 70
1.8 The site effect on decomposition rates of Schefflera octophylla in Nanjenshan forest, Taiwan ........................................................... 71
1.9 The effects of litterbag mesh size on decomposition rates on Schefflera octophylla in Nanjenshan forest, Taiwan ..................................... 72
1.10 Relationships between the decay rates of Schefflera octophylla and the densities of total invertebrates for three mesh size litterbags from Nanjenshan forest, Taiwan ............................................................ 73
2.1 Composition of litter amphibians in the creek and the windward sites of Nanjenshan forest, Taiwan ........................................................... 74
2.2 Abundance of juvenile and adult anurans on forest floor of Nanjenshan during the experimental period in 2001 ....................................... 75
3.1 Density of total invertebrates from the control and amphibian-excluded enclosures in Nanjenshan forest, Taiwan. .................................... 76
3.2 Multi-dimensional scaling (MDS) plot of invertebrate communities in litterbags from the control and amphibian-excluded enclosures at different time in Nanjenshan forest, Taiwan .............................. 77
3.3 The predatory effect of Bufo bankorensis on decomposition rates of Schefflera octophylla in Nanjenshan forest, Taiwan .................. 78
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